W ith 350,000 known

species, and scientific

estimates that mil-

lions more have yet to be identi-

fied, beetles are one of the most

remarkable and varied creatures

on earth. They range from the de-

lightful summer firefly to the one-

hundred-gram Goliath beetle.

Beetles offer a dazzling array of

shapes, sizes, and colors that en-

tice scientists and collectors across

the globe.

This collection covers six hundred

significant beetle species. Each

features a distribution map, basic

biology, conservation status, and

information on cultural and eco-

nomic significance. Full-color

photos show the beetles both at

actual size and enlarged to show

details. Based in the most up-to-

date science and accessibly writ-

ten, the descriptive text will ap-

peal to researchers and armchair

coleopterists alike.

Publication date

October 15, 2014

$55.00, cloth

978-0-226-08275-2

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2400 color plates

7 1/8 x 10 1/2

For a review copy or other

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contact:

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The Book of Beetles A Life-Size Guide to

Six Hundred of Nature's Gems

Edited by Patrice Bouchard

Contributors:

PATRICE BOUCHARD, YVES BOUSQUET, CHRISTOPHER CARLTON, MARIA LOURDES CHAMORRO,

HERMES E. ESCALONA, ARTHUR V. EVANS, ALEXANDER S. KONSTANTINOV, RICHARD A. B. LESCHEN,

STÉPHANE LE TIRANT, AND STEVEN W. LINGAFELTER.

THE BOOK OF BEETLES

THE UNIVERSITY OF C HICAGO PRESS

Chicago

THE BOOK OF BEETLESA L I F E - S I Z E G U I D E T O S I X H U N D R E D

O F N A T U R E ’ S G E M S

E D I T O R PAT R I C E B O U C H A R D

C O N T R I B U T O R S PAT R I C E B O U C H A R D , Y V E S B O U S Q U E T, C H R I S T O P H E R C A R L T O N ,

M A R I A L O U R D E S C H A M O R R O , H E R M E S E . E S C A L O N A , A R T H U R V. E VA N S , A L E X A N D E R K O N S TA N T I N O V, R I C H A R D A . B . L E S C H E N ,

S T É P H A N E L E T I R A N T, S T E V E N W. L I N G A F E L T E R

The University of Chicago Press, Chicago 60637The University of Chicago Press, Ltd., London© 2014 by Ivy Press LimitedAll rights reserved. Published 2014.Printed in China

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ISBN-13: 978-0-226-08275-2 (cloth)ISBN-13: 978-0-226-08289-9 (e-book)DOI: 10.7208/chicago/9780226082899.001.0001

Portions of this work were written and prepared by o�cers and/or employees of the U.S. Government as part of their o�cial duties and are not copyrightable.

Library of Congress Cataloging-in-Publication Data

The book of beetles : a life-size guide to 600 of nature ’s gems / edited by Patrice Bouchard ; with contributions by Patrice Bouchard, Yves Bousquet, Christopher Carlton, Maria Lourdes Chamorro, Hermes E. Escalona, Arthur V. Evans, Alexander Konstantinov, Richard A. B. Leschen, Stéphane Le Tirant, and Steven W. Lingafelter. pages : illustrations ; cm Includes bibliographical references and index. ISBN 978-0-226-08275-2 (cloth : alk. paper) — ISBN 978-0-226-08289-9 (e-book) 1. Beetles—Identification. 2. Beetles—Pictorial works. I. Bouchard, Patrice, 1973– editor. QL575.B66 2014 595.76—dc23

2014010716

∞ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).

Color origination by Ivy Press Reprographics

This book was conceived, designed, and produced by Ivy Press210 High Street, LewesEast Sussex BN7 2NSUnited Kingdomwww.ivy-group.co.uk

Creative Director PETER BRIDGEWATERPublisher SUSAN KELLYArt Director MICHAEL WHITEHEADEditorial Director TOM KITCHSenior Project Editor CAROLINE EARLECommissioning Editor KATE SHANAHANDesigner GINNY ZEALIllustrator SANDRA POND

LITHOCASE IMAGES © Jason Bond and Trip Lamb: Onymacris bicolor.

© Lech Borowiec: Cetonia aurata, Dytiscus marginalis, Lytta vesicatoria, Pachylister inaequalis, Pachnephorus tessellates

Karolyn Darrow © The Smithsonian Institution: Tetracha carolina

Anthony Davies © Her Majesty the Queen in Right of Canada as represented by the Minister of Agriculture and Agri-Food: Actinus imperialis, Anthrenus museorum, Brachycerus ornatus, Byctiscus rugosus, Erotylus onagga, Hister quadrinotatus quadrinotatus, Saprinus cyaneus, Spilopyra sumptuosa

Henri Goulet © Her Majesty the Queen in Right of Canada as represented by the Minister of Agriculture and Agri-Food: Calosoma sycophanta

© René Limoges: Chalcosoma atlas

© Kirill Makarov: Byturus tomentosus

© Udo Schmidt: Paranaleptes reticulata

© Maxim Smirnov: Lamprima adolphinae, Mormolyce phyllodes, Sulcophaneus imperator

© Chris Wirth: Alaus zunianus, Amblysterna natalensis, Calodema regalis, Thrincopyge alacris

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CONTENTSIntroduction 6

What is a beetle? 10

Beetle classi�cation 16

Evolution & diversity 18

Communication, reproduction & development 20

Defense 22

Feeding behavior 24

Beetle conservation 26

Beetles & society 28

The beetles 30

ARCHOSTEMATA 32

MYXOPHAGA 42

ADEPHAGA 48

POLYPHAGA 112

Appendices 640

Glossary 642

Classi�cation of the Coleoptera 646

Resources 648

Notes on contributors 650

Index of species and families 652

Acknowledgments 656

PATRICE BOUCHARD is a research scientist and curator of Coleoptera at the Canadian National Collection of Insects, Arachnids, and Nematodes.YVES BOUSQUET is a research scientist at the Canadian National Collection of Insects, Arachnids, and Nematodes.CHRISTOPHER CARLTON is a research scientist and director of the Louisiana State Arthropod MuseumMARIA LOURDES CHAMORRO is a research entomologist with the Systematic Entomology Laboratory at the National Museum of Natural History in Washington, D.C.HERMES E. ESCALONA is a visiting scientist at the Australian National Insect Collection-CSIRO.ARTHUR V. EVANS is a research associate at the Smithsonian Institution.ALEXANDER KONSTANTINOV is a research entomologist with the Systematic Entomology Laboratory at the National Museum of Natural History in Washington, D.C.RICHARD A. B. LESCHEN is a researcher at Landcare Research, New Zealand Arthropod Collection.STÉPHANE LE TIRANT is curator of the Montreal Insectarium.STEVEN W. LINGAFELTER is a research entomologist with the Systematic Entomology Laboratory, United States Department of Agriculture

INTRODUCTION

6

Beetles of the order Coleoptera, with nearly 400,000 described species, comprise one of the most diverse and important groups of animals on Earth. As such, coleopterists, biologists who specialize in the study of beetles, have a view of the natural world with a degree of resolution that is seldom seen through the study of other organisms.

One out of every �ve species of plants and animals is a beetle. Despite their riot of forms, colors, patterns, and behaviors, all beetles share a select suite of physical attributes, the most conspicuous of which are the leathery

INTRODUCTION

“From the small size of insects, we are apt to undervalue their appearance. If we could imagine a male Chalcosoma with its polished, bronzed coat of mail, and vast complex horns, magni�ed to the size of a horse or even of a dog, it would be one of the most imposing animals in the world.”CHARLES DARWIN, THE DESCENT OF MAN, & SELECTION IN RELATION TO SEX, 1871

ABOVE Beetles represent an exceptionally diverse group of organisms. Their importance for agriculture, forestry, culture, and science, and endless variations in structures (e.g., the male Atlas Beetle, Chalcosoma atlas) and microhabitat adaptations, have captivated humans for centuries.

ABOVE Some beetles feed on a single plant species; others feed on a wide range of hosts. Adults of the North American pest the Japanese Beetle (Popillia japonica) have been observed feeding on 300 hosts in approximately 80 plant families.

BELOW Most beetles are winged, but some have reduced hind wings and cannot �y, such as species in the African weevil genus Brachycerus.

or hardened forewings, or elytra (singular elytron). Depending on the species, elytra can help stabilize beetles in flight, protect their delicate hind wings and internal organs, conserve precious bodily fluids, capture bubbles of air underwater, and insulate them from extreme temperatures. Combined with their small and compact bodies and numer ous other morphological and behavioral adaptations, beetles exploit and thrive in niches unoccupied or underutilized by other animals in widely diverse terrestrial and freshwater habitats.

Although the sheer number of species prevents all but the most common or economically important beetles from having a meaningful and widely accepted common name, each known species does have a scienti�c name consisting of a genus (plural genera) and species (singular and plural) epithet that is universally recognized. To manage information e�ectively, coleopterists file each species into a nested system of hierarchical groups, or taxa (singular taxon), based on their shared evolutionary characteristics. Species is the most exclusive taxon, while the order Coleoptera is the most inclusive of beetle taxa.

Beetles communicate with one another through physical, chemical, or visual means, usually to locate a mate. Although most species engage in sexual reproduction, a few reproduce asexually by cloning themselves, a process known as parthenogenesis. Among beetles, limited parental care of the young is the exception, not the rule. The larvae and adults eat a variety of organisms, living and dead, especially plants. Those that prefer leaves, �owers, fruits, needles, cones, and roots can in�ict serious damage to food stores, gardens, crops, and managed timber. Some carnivorous beetles are used as biological control agents against agricultural or forestry pests, while scavenger species provide an essential service to clean study skeletons in natural history collections around the globe. Recently, the study of beetle structure and function has inspired scientists and engineers

9

INTRODUCTION

8

Physically impressive Large, colorful, horned, or exaggerated or unusually developed legs or mouthparts—species with special structures involved in speci�c behavior that has evolved through natural selection over millions of years and thus carries interesting underlying genetic information.

Each species is depicted by a razor-sharp life-size photograph and accompanied by a summary of its known distribution, habitat, and feeding habits. The map o�ers an indication of its global distribution, while the engravings afford readers another point of view. The scientific and common (if any) names are also provided, followed by the species’ author and year of description. Concise narratives covering natural history and related species are followed by a brief species diagnosis.

BEETLE COLLECTIONSBeetle collections carefully assembled over decades by thousands of dedicated professionals and amateurs provide essential data needed to identify and map sensitive species within a historical context. These scienti�cally valuable collections also serve as important resources for other scienti�c and educational endeavors, such as providing the specimens photographed for this book.

working in the rapidly growing �eld of biomimetics to develop and design new materials and products ranging from iridescent car paint and reusable adhesive tape to monetary security systems.

The Book of Beetles o�ers a glimpse of this incredible diversity through an overview of 600 beetle species presented within a framework based on their evolutionary relationships. The diversity of beetles is divided by species into four main chapters of the suborders. Within each section the arrangement is taxonomic, by family and then subfamily, and then within each subfamily in scienti�c name (genus, species) order.

SELECTION CRITERIAThe species were selected on the basis of several criteria, o�ering a unique taxonomic survey of the majority of beetle families from around the world:Scienti�cally compelling Subjects of focused scienti�c research or medicinal use, or inspirations for biomimetic and technological innovation. Curious natural histories Unusual adaptations, ability to live in extreme habitats, interesting symbioses, or engaging behaviors.Culturally signi�cant Mythological and religious symbols, uses in folk medicine, or entomophagy.Economically important Pest species, use as biological control agents, sources of products and services, or with a role in forensic entomology.Conservation Rare and threatened species.

ABOVE One of the most challenging tasks for this project was to select the 600 candidate beetles to cover from the overwhelming number of species. While some species such as the Agreeable Caterpillar Hunter (Calosoma sycophanta) (left) are visually attractive and well known biologically, others such as the Frog-legged Leaf Beetle (Sagra buqueti) (right) have notable unusual adaptations yet little is known about their biology. We hope that this book will stimulate the discovery and publication of new knowledge on beetles from around the world.

BELOW Millions of insects, including beetles, are preserved in natural history collections throughout the world. Specimens are typically dried and pinned, with a label attached to the same pin that includes important information such as the locality where the individual was encountered, the date it was collected, and the person who found it. Specimens are then placed in boxes or drawers, such as this one from the Alfred Russel Wallace (1823–1913) collection in the Natural History Museum, London, in order to protect them from museum pests and improve long-term preservation.

11

WHAT IS A BEETLE?

10

secretions or microscopic networks of cracks (alutaceous) resembling that of human skin. The surface is variously festooned with spines, hairlike setae, or flattened setae called scales, and sculpted with tiny bumps (tubercles), pit-like punctures, ridges, grooves (striae), or rows of punctures.

COLORThe colors of beetles are derived either from chemical pigments obtained from their food or structural properties of the outer layers of the exoskeleton. Most beetles are black as a result of melanin deposition during sclerotization, the chemical hardening process of the exoskeleton that occurs after emergence from the pupa, or eclosion. Microscopic surface sculpturing also influences beetle colors, as do patterns of setae, scales, or waxy secretions. Black desert darkling beetles (Tenebrionidae) are sometimes partially or completely covered with a white, yellow, or bluish-gray waxy bloom that re�ects light and helps to keep the beetle cool.

The brilliant iridescent and metallic colors of beetles are created by multiple re�ective layers in the exoskelton and scales, or a layer of highly complex photonic crystals that re�ect light at di�erent wavelengths to create speci�c metallic colors and shimmering iridescence. These structures are determined genetically, but their �nal form in individual beetles is determined by conditions experienced during growth and development.

WHAT IS A BEETLE?The English word “beetle” comes from the Middle English bityl or betyll and the Old English bitula, all of which mean “little biter.” Other commonly used names, such as “weevil” and “chafer” derived from Old English and Old High German, also relate to biting. Coleoptera, �rst coined by Aristotle in the fourth century bce and later adopted as an order of insects by Carl Linnaeus in 1758, is derived from the Greek words koleos, meaning “sheath,” and pteron, or “winged,” and was inspired by the tough elytra of beetles.

DISTINGUISHING FEATURESAmong other adaptations, beetles are distinguished from other insects by their chewing mouthparts, the conversion of their forewings into hardened elytra, their hind wings that fold lengthwise and across beneath the elytra, and their holometabolous development. Holometabolous insects pass through four very distinct life stages: egg, larva, pupa, and adult. The larvae and adults frequently di�er in habits and habitat, functioning in the environment as if they were two separate species.

Beetles, like other insects, crustaceans, arachnids, millipedes, centipedes, and their kin with segmented exoskeletons and jointed appendages (antennae, mouthparts, legs), are classi�ed in the phylum Arthropoda. Light and durable, the beetle exoskeleton is incredibly tough and rigid or characteristically soft and pliable, and provides protection and support. It serves as a platform for important tactile and chemosensory structures externally, while providing an internal framework that supports muscles and organs. The exoskeletal surface is smooth and shiny, or dulled by waxy

ABOVE The study of a superdiverse group of organisms such as beetles requires the close examination of a wide range of anatomical structures in order to recognize and classify species and larger groups correctly. Using clear and consistent terms for these structures is essential to clearly communicate results amongst those who are specialists and non-specialists.

RIGHT The forewings of beetles are uniquely modi�ed into hardened, non-foldable elytra that meet together in a straight line over the body when at rest. The elytra partly or completely cover the abdomen and can be opened to deploy the hind wings, as in this Lucanus cervus individual.

tarsomeres

coxa

trochanter

VENTRAL VIEWDORSAL VIEW

antenna

compound eye

pronotum

scutellumelytron

tibia

elytralsuture

tarsal claw

tarsus

mandible

labrum

front leg

middle leg

hind leg

mentum

prosternal process

elytral epipleuron

femur

abdominal ventrites

166

FAMILY

SUBFAMILY

DISTRIBUTION

MACROHABITAT

MICROHABITAT

FEEDING HABITS

NOTE

167

FAMILY

SUBFAMILY

DISTRIBUTION

MACROHABITAT

MICROHABITAT

FEEDING HABITS

NOTE

POLYPHAGAPOLYPHAGA

Xenodusa re�exa is a comparatively large, mahogany brown member of the rove beetle tribe Lomechusini. The broad, re�exed pronotum and unusual trichome bundles along the abdomen are characteristic of the genus. Similar glandular trichomes are found on various parts of the bodies of other myrmecophilous beetles, and presumably also function in producing and distributing glandular appeasement chemicals.

Adults and larvae of Xenodusa reflexa are adapted to an obligate association with formicine ants and spend most of their lives in ant nests. Specialized glandular setae on the adult

abdomen produce appeasement chemicals that are attractive to host ants. Both adults and larvae are fed by regurgitation by the ants (trophallaxis). They are considered nest

parasites because beetle larvae compete with ant larvae for care and feeding by the ants. Ants in the genus Camponotus

are the most frequently documented hosts for this species, but other species breed in Formica nests and overwinter in Camponotus nests.

RELATED SPECIES Four species of the genus Xenodusa occur in USA and

southern Canada, and an additional species occurs in Mexico. Xenodusa reflexa is the most widely distributed species. It differs from species that overlap in ranges by its relatively

larger size (3⁄16 in, or 4.5 mm) and hairy ventral surface. A widespread eastern species, X. cava, is similar, but lacks long hairs on the ventral surface of the body.

XENODUSA REFLEXA

XENODUSA REFLEXA (WALKER, 1866)

ADULT LENGTH

Adult length: 3∕16–¼ in (5.3–6.4 mm)

StaphylinidaeAleocharinaeNearctic: North America, north of MexicoForestsNests of formicine antsAdults and larvae are fed by hosts through trophallaxisA specialized nest parasite of formicine ants

The Long-necked Shining Fungus Beetle is unique among rove beetles and perhaps all beetles in possessing an extremely long neck that may be twice the length of the body in some male specimens. The neck is longer on average in males but its length is variable in both sexes.

From the head back, this remarkable insect looks like a typical member of the tribe Scaphidiini of the subfamily Scaphidiinae, but the extremely elongate neck in both sexes render it unique among rove beetles. The biology of the species is poorly known, but other members of the subfamily graze on �eshy or encrusting fungi on dead wood or other organic substrates in forest habitats as both adults and larvae. Adults may be encountered on exposed fungal surfaces day or night. They are wary and prone to flight or dropping, requiring a slow, stealthy approach if observations or photographic e�orts are to be successful.

RELATED SPECIES This species is placed in the same tribe as the enormous genus Scaphidium based on recent phylogenetic studies and general appearance, exclusive of the outsized neck. It is not likely to be confused with any other animal within its range, but some African brentid weevils have elongate forebodies comprising parts of the thorax and neck that are super�cially similar.

DIATELIUM WALLACEI

LONG-NECKED SHINING FUNGUS BEETLE PASCOE, 1863

ADULT LENGTH

½–¾ in (13–20 mm)

StaphylinidaeScaphidiinaeOriental: Sumatra and Sulawesi, Indonesia; Sarawak, MalaysiaTropical forestsFungus on logs and standing dead treesFungivorousThe neck may be twice the body length

Actual size

Actual size

182

FAMILY

SUBFAMILY

DISTRIBUTION

MACROHABITAT

MICROHABITAT

FEEDING HABITS

NOTE

183

FAMILY

SUBFAMILY

DISTRIBUTION

MACROHABITAT

MICROHABITAT

FEEDING HABITS

NOTE

POLYPHAGAPOLYPHAGA

Amblyopinodes piceus is an elongate, somewhat �attened, somber-colored rove beetle. The head in particular is unusually �attened and the mandibles are adapted to grasping mammal hairs. Elongate, movable hairs on the underside of the abdomen are unique to the genus.

Members of this species and others in the tribe Staphylinini, subtribe Amblyopinina are unique among rove beetles in their adaptations to life on the bodies and in the nests of small mammals, mainly rodents. Adults of this species grasp the hair and skin between the ears of South American Water Rats (Nectomys squamipes) and travel with the animal, and the larvae occur in water rat nests. Early entomologists thought these beetles were parasitic on the animals, but later researchers revealed that they are predators of �eas and serve a bene�cial function for the rats by reducing �ea populations in their nests. The flattened morphology of the beetle ’s head is similar to that of other unrelated species that live on mammals.

RELATED SPECIES At least five other species of the genus occur in the same region. Identi�cation of species is based on examination of internal male sex structures and, to a lesser extent, the arrangement of specialized hairs on the body. Other genera of the tribe are similar and may be distinguished using available keys and by mammal host associations.

AMBLYOPINODES PICEUS

AMBLYOPINODES PICEUS(BRÈTHES, 1926)

ADULT LENGTH

¼–5/16 in (6.5–7.5 mm)

StaphylinidaeStaphylininaeNeotropical: central South AmericaForests near rivers and streamsThe body and nests of South American Water Rats (Nectomys squamipes) Predatory on �easPhoretic on small mammals and found in their nests feeding on �eas

Actinus imperialis is unusually large for a staphylinid beetle, with a metallic green head and pronotum and brilliant metallic purple elytra. The abdomen possesses a triangular orange spot near the posterior end. The punctures on the head are diagnostic for this species.

Adults of this unusually large and gaudily colored rove beetle frequent smelly substrates such as carrion and dung that are likely to attract flies. Adults actively seek

out and destroy predators that feed on maggots and opportunistically capture adult flies. The immature stages presumably share the predatory habits of the adults but details of their biology are not well documented. One source mentions an association with cardamom, without giving additional details. Adults exhibit strong sexual dimorphism of the mandibles and head, with males often having larger heads and longer mandibles than females.

RELATED SPECIES Two similar species are known from the genus, with Actinus macleayi from northern Australia di�ering mainly in having a less strongly punctured head and pronotum, in addition to other minor differences. In other respects A. imperialis super�cially resembles other large members of the staphylinid subtribe Philonthina, but few are as brightly colored.

ACTINUS IMPERIALIS

ACTINUS IMPERIALISFAUVEL, 1878

ADULT LENGTH

5/8–7/8 in (16–22 mm)

StaphylinidaeStaphylininaeAustralian: New Guinea and extreme northern AustraliaTropical forestsLeaf litter, dung, carrionPredatory on �iesPreys on �ies associated with animal carcasses

Actual size

Actual size

208 209

POLYPHAGAPOLYPHAGA

Sulcophanaeus imperator is a large and bulky beetle that is dull to weakly shining black mixed with brilliant metallic greens, golds, and blues, and coppery red re�ections. Males have a distinct backward-pointing horn on the head, while the females lack this armature. Both males and females have front tarsi.

Sulcophanaeus imperator adults reach their peak of activity from January to March. They �y during the day in search of fresh feces of humans and various domesticated animals, and can be common in cattle pastures. They typically work in pairs to dig their nests directly beside or underneath animal droppings. Males push plugs of dung down the tunnel to females that fashion them into brood balls within the brood chamber. Dung burial reduces the loss of pasturage due to the growth of rank herbage, releases nutrients back to the soil, and disrupts the life cycles of pest organisms developing in the feces.

RELATED SPECIESThe genus Sulcophanaeus consists of 14 species, all but four of which occur in South America. Three of the latter species inhabit Central America, including an endemic, while the fourth species is endemic to Jamaica. With its green, gold, and red color forms, Sulcophanaeus imperator is by far the most colorful and variable species in the genus.

SULCOPHANEUS IMPERATOR

SULCOPHANEUS IMPERATOR (CHEVROLAT, 1844)

ScarabaeidaeScarabaeinaeNeotropical: northern Argentina, southern Bolivia, western ParaguayDry forest, dry thorn forest, and pastureOften found in pastures in cattle-raising areasAdults commonly found in cattle dungThis is the most colorful and variable species of Sulcophanaeus

The Sacred Scarab Beetle has a broad, smooth patch and narrow groove along the posterior margin of the pronotum. The middle tibiae each have two oblique rows of short setae. The tips of the posterior tibiae are prolonged, forming a narrow plate beneath the tarsus. Males have a reddish fringe of setae along the inner margins of their hind tibiae.

Scarabaeus sacer uses its rakelike forelegs to fashion balls from fresh dung, then lays an egg inside the ball before rolling it away and burying it. The grub feeds and completes its development inside the ball. These beetles were revered in ancient Egypt as symbols of Khepri, a manifestation of the sun god Ra, because their dung-rolling activities were considered symbols of the forces that moved the sun, represented by the dung ball, across the sky. The species was also associated with rebirth, and the beetles, along with their likenesses carved in

stone, were frequently buried with the dead.

RELATED SPECIESScarabaeus contains 139 species in four subgenera that inhabit Afrotropical, Palearctic, and Oriental regions. These small to relatively large dung beetles have four distinct teeth across the clypeus, anterior coxae and

femora that are not enlarged, and lack front tarsi. Scarabaeus sacer is distinguished from other Palearctic species, in part, by

the �nely notched posterior pronotal margin, and the features of the middle and hind tibiae.

SCARABAEUS SACER

SACRED SCARAB BEETLELINNAEUS, 1758

ADULT LENGTH

11∕32–19∕16 in (26–40 mm)

ScarabaeidaeScarabaeinaePalearctic: southern and central Europe, North Africa, Middle East, portions of AsiaSteppe, forest-steppe, and semidesert In the vicinity of accumulations of fresh dungAdults strain nutrients from dung; larvae eat solid wasteRegarded as a sacred symbol by the ancient Egyptians

ADULT LENGTH

1 1/16–11/8 in (18–28 mm)

FAMILY

SUBFAMILY

DISTRIBUTION

MACROHABITAT

MICROHABITAT

FEEDING HABITS

NOTE

FAMILY

SUBFAMILY

DISTRIBUTION

MACROHABITAT

MICROHABITAT

FEEDING HABITS

NOTE

Actual size

Actual size

492

FAMILY

SUBFAMILY

DISTRIBUTION

MACROHABITAT

MICROHABITAT

FEEDING HABITS

NOTE

493

FAMILY

SUBFAMILY

DISTRIBUTION

MACROHABITAT

MICROHABITAT

FEEDING HABITS

NOTE

POLYPHAGAPOLYPHAGA

Strongylium auratum is an elongated beetle with bright metallic green to reddish-purple re�ections on the dorsal surface and legs. Each elytron has nine distinct rows of transverse punctures. Antennae are rather long, reaching beyond the base of the pronotum. Antennomeres are slightly wider at their apex and are speckled with small circular white sensory structures. The last two abdominal ventrites are contrasting yellow-reddish in color.

Based on the abundance of darkling beetles in most dry environments on the planet, many people assume that all species in this family are dark brown to black and are ground-dwellers. This is clearly not the case for species in the tribe Stenochiini, which includes the colorful forest-dwelling genera Stronglyium and Cuphotes. Strongylium is certainly one of the most species-rich genera in the family, with nearly 1,000 described species and many more undescribed, especially in tropical areas. Strongylium auratum is relatively abundant in Neotropical forests and can be encountered at elevations of 5,000 ft (1,500 m) and more.

RELATED SPECIES The overwhelming diversity in Strongylium, coupled with an almost complete lack of comparative studies of its species, is a major impediment to taxonomic and biological studies. New species continue to be described every year, however, primarily based on di�erences in color patterns, sculpture of their cuticle, and sexual characters. Although most species are elongated and have well-developed �ight wings, such as S. auratum, others are �ightless and more convex.

ADULT LENGTH

½–¾ in (12–19 mm)

STRONGYLIUM AURATUM

STRONGYLIUM AURATUM(LAPORTE, 1840)

TenebrionidaeStenochiinaeNeotropical: from Mexico south to BoliviaTropical forestsAdults live on the surface of trees or under bark, and larvae in woodImmature stages probably develop in decaying woodOne of the most diverse and colorful genera in the family Tenebrionidae

Frégate Island Giant Tenebrionid Beetle adults are pale gray to dark brown and have broadly rounded elytra in dorsal view, which are covered by a small number of apically rounded and shiny tubercles. The elytra are completely fused along the midline, and �ight wings are absent. The legs are relatively long and males have curved tibiae.

The Frégate Island Giant Tenebrionid Beetle is one of 12 beetle species ranked Critically Endangered on the International Union for Conservation of Nature’s Red List of Threatened Species (see also the Delta Ground Beetle, Elaphrus viridis, and American Burying Beetle, Nicrophorus americanus), and is restricted to Frégate Island in the Seychelles. The accidental introduction of Brown Rats (Rattus norvegicus) to the island in 1995 nearly caused the extinction of the flightless beetle, along with the endemic Seychelles Magpie-robin (Copsychus sechellarum), before the rodents were eradicated in the early 2000s.

RELATED SPECIES Flightless tenebrionid genera were previously grouped

together on the basis of the absence of flight wings and their sealed elytra. Recent studies on internal organ systems such as the defense glands and the female genital tube now suggest that the older

classi�cation did not re�ect natural groupings. Although the monotypic genus Polposipus belongs to the diverse

subfamily Stenochiinae based on internal characters, its closest relatives have not yet been established.

ADULT LENGTH

1–13/16 in (25–30 mm)

POLPOSIPUS HERCULEANUS

FRÉGATE ISLAND GIANT TENEBRIONID BEETLE

SOLIER, 1848

TenebrionidaeStenochiinaeEndemic to Frégate Island, Seychelles Tropical forestAdults and larvae are most commonly associated with Pterocarpus indicus Decaying wood and barkKnown only on Frégate Island, a 0.8 sq miles (2 sq km) island in the Indian Ocean

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595

POLYPHAGAPOLYPHAGA

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Eugnamptus nigriventris is clothed in �ne, erect setae and has a reddish head, pronotum, and legs, and bluish-green elytra. In the male, the length of the rostrum is shorter than the length of the head and the antennae are inserted close to the apex. In the female, the rostrum is longer than the head, and the antennae are inserted near the middle. The head and pronotum are narrower than the base of the elytra.

This species exhibits maternal care by individually placing each egg between the epidermal layers of a dead leaf of the host tree (oaks, Quercus spp.). In spring, a female uses her mandibles to cut an oviposition scar on a leaf that dropped to the ground the previous fall. She then oviposits a single egg into a cavity she created between the upper and lower epidermis and seals the epidermal tissue by pinching with her mandibles. The larva completes its entire development by feeding on the epidermal tissues of the dead leaf. Pupation takes place in the soil and adults emerge the following spring.

RELATED SPECIES There are approximately 100 species included in Eugnamptus. Eugnamptus is similar to Hemilypus, Acritorrhynchites, and Essodius, but can be distinguished by characters of the rostrum, distance between the eyes, length of basitarsal segments, prominence of the �rst abdominal suture, extent of dorsal punctation, and elytral length as it relates to the pygidium.

EUGNAMPTUS NIGRIVENTRIS

EUGNAMPTUS NIGRIVENTRIS(SCHAEFFER, 1905)

ADULT LENGTH

¼ in (5.7–6.3 mm)

AttelabidaeRhynchitinaeNearctic and Neotropical: USA (Arizona, New Mexico, Texas), Mexico (Chihuahua, Sonora, Oaxaca)Oak forestsFoliage, leaf litterEpidermal tissue of Quercus spp. The larvae are leaf-miners

Byctiscus rugosus is a bright metallic, shiny green weevil with reddish re�ections on the head and legs. The elytra are covered with prominently punctate striae. The pronotum is narrower than the base of the elytra, which are quadrate. The head is narrow and the rostrum is almost twice as long as the head. The antennae are not elbowed and emerge near the apex of the rostrum; the last three antennomeres are almost twice as wide as the preceding ones.

Ritualized contests and aggressive �ghting behavior are known to occur between males of species in this genus. The males extend their forelegs outward

while rearing up on their mid and hind legs, grabbing each other with their forelegs and touching their rostra.

The elongate tarsal setae may help enhance visual displays of this aggression. Females of the charismatic Byctiscus rugosus roll leaves into complicated cylindrical tubes for the reception of the egg. The larva feeds within the tube, where it completes its development.

RELATED SPECIES Byctiscus includes 27 species from the Palearctic and Oriental

regions. Species are classi�ed into two subgenera: Byctiscus and Aspidobyctiscus. The tribe Byctiscini has an exclusively

Old World distribution and includes 12 genera in two subtribes. The related Pear Leaf-roller Weevil

(B. betulae) is known to cause damage to grapevines, pears, and other broadleaved trees and shrubs.

ADULT LENGTH

¼–5/16 in (6.5–8 mm)

AttelabidaeRhynchitinaePalearctic: China, Japan, Kazakhstan, Korea, Mongolia, RussiaForestLeaf rolls, vegetationHost plants include species of Malus, Pyrus, Sorbaria, and Populus The males are known for their wrestling behavior

BYCTISCUS RUGOSUS

BYCTISCUS RUGOSUS(GEBLER, 1830)

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Brentus anchorago is a greatly elongate black weevil with reddish-orange vittae on the elytra. It is one of the largest weevil species in North America. Males are strikingly longer, with the long, slender prothorax narrowing in the middle and the elongate, narrow rostrum about as long as the prothorax. Females have a tear-shaped prothorax, broadening basally, and a rostrum that is about half the size of the prothorax.

Most brentid species exhibit sexual dimorphism and Brentus anchorago is no exception: Some individuals may be up to �ve times larger than the smallest. Both sexes engage in combat, and those with longer bodies and rostra, which are used as weapons, are more successful in securing a mate. There is an overall preference by both sexes for larger mates, thus skewing populations toward larger-bodied individuals. Females chew holes into the decaying wood of primarily Gumbo-limbo (Bursera simaruba) trees to oviposit. Adults can be readily found in large numbers under the bark of dead logs.

RELATED SPECIES Brentus and Cephalobarus are currently classified in the Neotropical tribe Brentini. Thirty-seven species are included in Brentus. Of these, B. cylindrus has been reported from Polynesia (Marquesas, Tahiti), where it is possible that it was introduced. The �rst brentid species described by Linnaeus in 1758 were B. anchorago and B. dispar, originally under the genus Curculio.

BRENTUS ANCHORAGO

BRENTUS ANCHORAGO(LINNAEUS, 1758)

ADULT MALE LENGTH

3/8–11 5/16 in (9–50 mm)

ADULT FEMALE LENGTH

5/16–11/16 in (8–27 mm)

BrentidaeBrentinaeNearctic and Neotropical: USA (Florida) to ParaguayTropical and subtropicalDecaying wood; under barkAdults feed on sap or visit �owers for nectar; larvae bore into dead wood and possibly feed on sap or fungal myceliaThe species exhibits sexual dimorphism and is the largest weevil in North America

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Amorphocephala coronata is shiny reddish-brown beetle with a narrowly elongate body. Its most outstanding feature is its large, complex head, with the hind rostrum (just below the frons) deeply concave and bearing brushes of sti� setae. The head is sexually dimorphic, the males having a more robust front rostrum with large sickle-shaped mandibles, and the females having elongate, cylindrical mandibles.

This species is a facultative ant associate, usually of Camponotus but tolerated, after minor hostility, by other

ant groups. When introduced to a Camponotus colony, these brentids face aggression by the workers until the ants discover and begin to lick glandular secretions from pubescent areas on the beetle ’s head. Apparently, Amorphocephala coronata exhibits a pseudoaltruistic behavior, whereby it regurgitates to the colony part of the food received from host worker ants. Ants have been observed tending to these brentids and actively trying to retain them in their nest. The species is gregarious, with many individuals found together.

RELATED SPECIES This genus belongs in the tribe Eremoxenini (which is sometimes treated as the subtribe Eremoxenina within the Brentini), an almost exclusively myrmecophilous group. Other genera in this group include Cobalocephalus, Eremoxenus, and Symmorphocerus. Amorphocephala contains 20 species from the Palearctic and Afrotropical regions. The species can be separated mainly by characteristics of the head, prothorax and antennae.

AMORPHOCEPHALA CORONATA

AMORPHOCEPHALA CORONATA

(GERMAR, 1817)

ADULT LENGTH

3/8–1 1/16 in (9–18 mm)

BrentidaeBrentinaePalearctic: southern Europe, Algeria, Morocco, Israel, Syria, Iran, RussiaTemperate humid forests containing predominantly oaks (Quercus spp.)Nests of Camponotus, Lasius, Crematogaster, Pheidole, Tapinoma, and Myrmica antsUncertainMyrmecophilous; has an unusual head morphology

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